A TAXONOMIC REVISION OF
THE PAEONIA ANOMALACOMPLEX (PAEONIACEAE) 1

Hong De-Yuan 2 and Pan Kai-Yu 2

ABSTRACT
Based on field observation, examination of over 350 sheets of exsiccatae, and investigation of all the relevant types,
a taxonomic revision of the Paeonia anomala complex is presented. Typifications are clarified for P. anomala, P.
intermedia, P. hybrida, P. sinjiangensis, P. altaica, and P. veitchii. The identity of three well-known taxa, P. anomala,P. hybrida, and P. intermedia, is also clarified. Two species, P. anomala and P. intermedia, are recognized, with thecircumscription of P. anomala emended to include P. veitchii as its subspecies. Paeonia anomala includes two subspecies: subspecies anomala and subspecies veitchii with the former distributed in Central Asia, Siberia, and thenortheastern European part of Russia, while the latter occurs in China southeast of the Gobi Desert. Paeonia intermedia
is distributed in Central Asia, but with an isolated locality in Georgia. Paeonia sinjiangensis is treated as a synonymof P. anomala, and P. hybrida Pallas is treated as a synonym of P. tenuifolia L. The lectotypes of Paeonia laciniata,
Paeonia beresowskii, and Paeonia intermedia are designated here in the present paper.
Key words: Paeonia, Paeonia anomala complex, Paeonia hybrida, Paeonia intermedia, Paeoniaceae.

The Paeonia anomala L. complex (Paeoniaceae)
comprises a group of herbaceous peonies in Central
Asia, Siberia, and adjacent northeastern European
regions. They are characterized by leaves biternatewith leaflets decurrent at the base, leaflets finely
segmented, with segments of a lower (the best developed) leaf ranging from 70 to 100 in number and
4–32 mm in width, and by bristles along veins on
the upper blade surface. The only species withwhich it may be confused is Paeonia tenuifolia L.,
but the latter has leaves even more finely segmented, with segments more than 130 in number
and 0.5–6 mm in width.
Since the late 1970s, two new species in the
Paeonia anomala complex have been describedfrom Xinjiang, the Central Asian part of China: P.
sinjiangensis K. Y. Pan (1979) and P. altaica K. M.
Dai & T. H. Ying (1990). For a better understanding of the species group in Xinjiang, we made an
expedition in 1993 to the Altai and the Tienshan
in Xinjiang and found that morphological featuresof the group in Xinjiang were not consistent withthe descriptions by Schipczinsky (1921, 1937) andGamaulova (1961); Stern’s (1946) treatments andnomenclature are different from theirs but do notreflect the reality of the group in Xinjiang, either.
Thus, we realized that the P. anomala complex

needed a comprehensive taxonomic revision andreview of the nomenclature. For this purpose, thefirst author examined all available specimens of
this group in the herbaria at BM, HNWP, K, LE,
NWTC, PE, SHI, SHMU, TBI, XJBI, XJNU, andXJU (see Appendix 1). In addition, the types of P.
anomala, P. hybrida, P. intermedia, and the othersix specific and varietal names were examined at
LINN (Herbarium of the Linnean Society), BM, K,
and PE, respectively.
HISTORICAL REVIEW
Within the Paeonia anomala complex the typespecies, P. anomala, was described from Siberia by
Linnaeus (1771). Pallas (1789) described three additional species, P. laciniata and P. sibirica from
Siberia, and P. hybrida from a plant raised fromseeds of P. tenuifolia cultivated in the Botanic Garden of the St. Petersburg Academy in Russia.
Paeonia sibirica Pallas (1789) shares an illustration(tabula) with P. laciniata, but does not have a description. In his monograph of Paeonia, Anderson(1818) recognized only P. anomala, reducing P. hybrida as a synonym of P. tenuifolia for the first time.
De Candolle (1818) recognized P. anomala, P. hybrida, and P. laciniata, but later (1824) treated P.

1
The authors are grateful to the National Geographic Society for financial support (Grant 6408-99), which allowed
us to examine herbarium specimens at the Komarov Institute of Botany, the Russian Academy of Sciences, and to
conduct fieldwork in Georgia. The project is also supported by the National Natural Science Foundation of China (NSFC
Grant 30130030). We are grateful to the curators of the following herbaria: BM, HNWP, K, LE, NWTC, PE, SHI,SHMU, TBI, XJBI, XJNU, and XJU. We sincerely thank Zhou Shi-Liang for his assistance with fieldwork, and Li QiaoLing and Ma Li-Ming for their help in preparation of the manuscript.
2
Laboratory of Systematic and Evolutionary Botany, Institute of Botany, The Chinese Academy of Sciences, Xiangshan, Beijing 100093, China. hongdy@ns.ibcas.ac.cn.

ANN. MISSOURI BOT. GARD. 91: 87–98. 2004.

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laciniata as a synonym of P. anomala. Meyer(1830) described the fourth species, P. intermedia,
from the Altai Mountains. Ledebour (1842) cited P.
anomala, P. hybrida, and P. intermedia, treating P.
laciniata as a synonym of P. anomala. Trautvetter(1860) recognized only one species, reducing P. hybrida as a variety of P. anomala, and treating P.
intermedia as a form of variety hybrida. Lynch(1890) cited P. hybrida and P. anomala, and statedthat the former was native to the Caucasus. Huth(1892) adopted the widest species concept of P. anomala, which includes four varieties: var. typica,
var. hybrida (5 P. intermedia), var. nudicarpa, andvar. emodi (5 P. emodi, confined to the western
Himalayas). A further treatment by Krylov (1901)
divided the complex into two species, P. anomala
and P. hybrida, and reduced P. intermedia as a variety of the latter, P. hybrida var. intermedia (C. A.
Meyer) Krylov, in addition to the typical variety. In
the area of West Tienshan (Kirghizia and Xinjiang,
China), Trautvetter (1904) enumerated only P. anomala, treating P. intermedia as its subspecies.
Schipczinsky (1921) basically followed Krylov(1901), recognizing two species, P. anomala (withtwo varieties, var. anomala and var. nudicarpa) andP. hybrida (with two varieties, var. hybrida and var.
intermedia), and the same treatment was adopted
for the Flora of the USSR (Schipczinsky, 1937).
These two Paeonia species were grouped by him
as series 4, Dentatae Kom. (Schipczinsky, 1937: 33,‘‘leaf lobes incised or with dentate margin’’). Rootsof both species were described as tuberous. In contrast, Stern (1946) recognized only one species in
his monograph of Paeonia, P. anomala, in which
two varieties were recognized: variety anomala withglabrous carpels, and variety intermedia with tomentose carpels. He did not mention roots of thisgroup. Stern (1946: 113) considered P. hybrida Pallas as an ambiguous name: ‘‘It is doubtful what
species Pallas intended by this name and, as it is
not possible to discover what he meant, this nameof P. hybrida has been omitted.’’ However, in the
Flora of Kazakhstan, Gamaulova (1961), following
Krylov (1901) and Schipczinsky (1921, 1937), described two species, and still recognized P. hybrida
as a valid name.
While working on Paeonia for the Flora Reipublicae Popularis Sinicae, Pan (1979) found a specimen from Xinjiang whose roots were basipetally attenuate. According to Schipczinsky (1937) andGamaulova (1961), Paeonia anomala and P. hybrida have tuberous or fusiform roots, so the plant was
described by her as a new species, P. sinjiangensis
K. Y. Pan; the plants with tuberous or fusiform roots
were treated as P. anomala with two varieties: va-

riety anomala and variety intermedia (Pan, 1979).
Also from Xinjiang, Dai and Ying (1990) describedanother new species, P. altaica, which was statedto have flowers larger than P. sinjiangensis and one
or two underdeveloped flower buds in addition to
the terminal blooming flower.
Interestingly, Schmitt (1999) failed to recognize
the distinct differences between Paeonia anomala
and P. intermedia in the root and calyx, and thus
treated them as a single species, but still recognized P. sinjiangensis and P. altaica as distinct species.
From the above it is clear that Paeonia hybrida,
P. intermedia, and P. anomala have been variously
treated. With this paper we hope to clarify the biological attributes and variations of the P. anomala
complex.
OBSERVATIONS AND DISCUSSION
To answer how many species were really present
in Xinjiang and what biological features they have,the first author and coworkers (Hong et al., 1994)
made an expedition to the Tienshan and Altai
Mountains, conducting field observations and population samplings of Paeonia in 1993. Seven populations in total were observed, covering thosecharacter states previously described for the P. anomala complex (Table 1). We found there were only
two species in Xinjiang that differed distinctly fromeach other in the root and calyx. One species had
the roots basipetally attenuate (carrot-shaped) andsepals mostly caudate (Fig. 2), preferring relatively
moist habitats in woods. This was treated as P. sinjiangensis. The other had roots tuberous or fusiform
and sepals mostly (at least 2) rounded but not caudate at apex, growing on sunny, shrubby or grassy
slopes, or in sparse woods. This latter was recognized as P. anomala. We also found the presenceor absence of indumentum on the carpels in thisspecies group to be a polymorphism, i.e., individuals with carpels either glabrous or from sparsely
to densely pubescent could be found within a single
population (Table 1). The indumentum on carpels
was demonstrated as quite variable, although it wasconsidered by the previous authors as taxonomically valuable. Therefore, any taxonomic division
based on this character, such as that between P.
anomala and P. anomala var. nudicarpa (by Huth,
1892) or P. anomala and P. anomala var. intermedia (by Stern, 1946), is artificial. Also not observed by the previous authors was the clear differentiation in root and calyx in this group, which
are closely correlated with each other (Fig. 2).
We also found that in the form with carrot-shaped

5(5)
green
fusiform to tuberous (5)2(4); 3(1)0(1); 5(3)green
fusiform to tuberous (6)3(2); 4(1)0(2); 5(1)green
fusiform to tuberous (2)2(1); 3(2)5(4); 0(2)0(1); 2(1); 3(5)0(4); 1(1)2(1); 3(5)greencarrot-shaped (2)fusiform to tuberous (3)0(2)
pale green
carrot-shaped (3)0(2); 1(3)pale green
carrot-shaped (3)D. Y. Hong et al. 1

D. Y. Hong et al. 4

D. Y. Hong et al. 0125

T. H. Ying 1006–1022
D. Y. Hong et al. 2

D. Y. Hong et al. 5

Altai: Mt. Halamaryi, NW of Altay City, 1200 m, Populus forest by stream
Altai: Xiaodong Gou Valley, NE of Altay City, 1060m, Populus-Betula-Picea forest at valley bottom
Altai: Habahe County, 1200–1550 m, in Betula forests
Altai: Mt. Halamaryi NW of Altay City, 1300 m, S
slope, rocky Berberis-Spiraea bushes
Altai: Mt. Halamaryi, NW of Altay City, 1000 m,
sparse bushes
Altai: Xiaodong Gou, NE of Altay City, 1060 m, W
slope, rocky sparse Spiraea-Berberis bushes
Tienshan: Yining County, Yining Forest Farm, 1150–
1200 m, N slope, sparse Prunus-Malus forest with
Cotoneaster & Spiraea
D. Y. Hong et al. 3

Number of
non-caudate sepals
Leaf color
Roots
Locality and habitat
Population

0(1)Table 1. The populations sampled and their characters in the Paeonia anomala complex in Xinjiang, China (all the vouchers are in PE). The figures before parentheses indicate
character states, while those in parentheses indicate the number of individuals observed. The indumentum on carpels is divided more or less arbitrarily into six grades with ‘‘0’’expressing totally glabrous, ‘‘1’’ extremely sparsely hairy, ‘‘5’’ entirely covered with hairs, while ‘‘2,’’ ‘‘3,’’ and ‘‘4’’ are in between.

roots, the petals varied greatly in length, from 3.5
to 6.5 cm, and in the Altai Mountains some individuals of this form possessed one or two additional
underdeveloped flower buds similar to Paeonia
veitchii. This was again a polymorphism, and notconsidered by us to be a stable character, andtherefore P. altaica was reduced to a synonym of P.
sinjiangensis (Hong et al., 1994).
After examination of about 200 sheets of herbarium specimens in the Komarov Institute of Botany (LE) in 1999, the first author found that two
species could also be distinguished in this complex:
one with the roots carrot-shaped and sepals mostly
caudate, while the other with the roots tuberous or
fusiform, and sepals mostly non-caudate. These biological features match those on plants observed in
Xinjiang (Hong et al., 1994). Gamaulova (1961,
tab. 2) described roots of Paeonia anomala as fusiform-thickened, and those of P. hybrida as tuberous-thickened. However, roots of P. intermedia (5P. hybrida sensu Schipczinsky and Gamaulova)
vary in shape from tuberous consistently to fusiform, and this variation could be found within populations or even on the same individual, whereasin P. anomala neither fusiform-thickening nor tuberous-thickening were found.
Contradictions exist between our natural observations (Hong et al., 1994) and earlier descriptions
by Schipczinsky (1937) and Gamaulova (1961).
Stern’s (1946) treatment of this complex as one species with two varieties (P. anomala and P. anomala
var. intermedia) is also at variance. For a clear taxonomic revision and correct nomenclature, one remaining issue ought to be resolved: the identity ofPaeonia anomala, P. hybrida, and P. intermedia.
The type specimen of P. anomala, with two visible
sepals both caudate and relatively thin leaves, isperfectly consistent with the form with carrotshaped roots. According to the type specimen at
LINN [Siberia, no. 692. 3], field observation, andexamination of over 350 sheets of exsiccatae, P.
sinjiangensis and P. altaica should be treated assynonyms of P. anomala; the descriptions of rootsof P. anomala by Schipczinsky (1937) and Gamaulova (1961) are erroneous. The type of P. hybrida(at BM), on which Pallas’s (1789) tab. 86 was apparently based, has leaf segments 3–4 mm in width,and at least more than 100 in number, and is thus
within the range of variation in P. tenuifolia L.
(Hong & Zhou, 2003). Therefore, P. hybrida Pallas
should be treated as a synonym of P. tenuifolia, asAnderson (1818) suggested nearly 200 years ago.
The type specimen of P. intermedia at K possesses
tuberous roots, mostly non-caudate sepals, and relatively thick and narrow leaf segments. The plants0(2); 1(1); 5(1)Hong & Pan
Paeonia anomala Complex

Indumentum
on carpels

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that have tuberous or fusiform roots, are distributed
widely from the Altai to Tadzhikistan and Uzbekistan, and were named P. hybrida by Schipczinsky(1921, 1937) and Gamaulova (1961) correspond to
this type specimen very well.
Stern’s (1946) statement concerning the name ofPaeonia hybrida is not correct, confusing P. intermedia with P. anomala. Pan’s (1979) treatment of
the P. anomala complex (a new species, P. sinjiangensis, and P. anomala with var. intermedia) is also
unjustifiable. Although Hong and his coworkers(Hong et al., 1994) properly described two species
and clearly stated their morphological and ecological differences for this group, the nomenclature
they used is misapplied.
The Paeonia anomala complex comprises two
species: P. anomala and P. intermedia. Paeonia
emodi, which was treated as a variety of P. anomala
by Huth (1892), remains a distinct species, withleaves ternate, without bristles on the upper surface, leaflets much less segmented, segments nomore than 40 in number and 1.5–4 cm wide, carpels usually two, less frequently one, very rarely
three, and flowers white. Paeonia veitchii Lynch
was described from Sichuan Province, China(Lynch, 1890), and was said (Stern, 1946; Pan,
1979) to differ from the P. anomala group in having
several flowers instead of a single flower on a stem.
According to our observation, one or two underdeveloped flower buds sometimes also exist in addition to one normal and terminal flower in P. anomala, while P. veitchii sometimes has flowers
solitary (P. veitchii var. uniflora), although it has 3
or 4 flowers or 1 to 3 underdeveloped flower budsin addition to the terminal flower on a stem. Thus,
P. veitchii was reduced to a subspecies of P. anomala (Hong et al., 2001).

Paeonia laciniata Pallas, Fl. Ross. 1(2): 93, tab. 85, subP. sibirica, 1789. TYPE: the illustration (tab.) citedin the protologue (lectotype, designated here!).
Paeonia sibirica Pallas, Fl. Ross. 1(2): tab. 85. 1789.

KEY TO THE PAEONIA ANOMALA COMPLEX
1a. Roots carrot-shaped, never tuberous or fusiform;
sepals all or mostly caudate, less frequently one
or very rarely two non-caudate --------- 1. P. anomala
2a. Flowers solitary, rarely 1 or 2 underdeveloped flower buds present in addition to terminal flower on a stem ------------------------------------------------------ 1a. P. anomala subsp. anomala
2b. Flowers usually 2 to 4 on a stem or 1 to 3
underdeveloped flower buds present in addition to terminal flower, rarely flowers solitary on a stem ---------------------------------------------------------------------- 1b. P. anomala subsp. veitchii
1b. Roots fusiform or tuberous; inner 2 to 4 sepals
rounded but non-caudate at apex 2. P. intermedia

TAXONOMIC TREATMENT
1. Paeonia anomala L., Mant. 2: 247. 1771.
TYPE: Siberia, no. 692.3 (holotype, LINN!).
Figure 1A.

Pallas’s (1789) illustration (tab.) 85 has only one
name, P. sibirica, and apparently he used this illustration also for P. laciniata because one sees‘‘Paeonia laciniata Tab. LXXXV’’ on p. 93. Paeonia laciniata was described in detail by Pallas, but
no description of P. sibirica was given by him.
Perennials; tap roots $ 50 cm long, carrotshaped, basipetally attenuate, up to 2 cm diam.,lateral roots slender, neither tuberous nor fusiform.
Leaves biternate; leaflets finely segmented; lower
leaves with segments 70 to 100 in number, 8–32
mm in width. Flowers solitary or 2 to 4 on a stem,
often only terminal one fully developed and blooming; sepals 3 to 5, mostly caudate at apex, rarely 1
or very occasionally 2 non-caudate; carpels 2 to 5,
from glabrous to densely tomentose.
The species prefers relatively moist habitats,growing in forests, on the edges of forests, or rarely
in bushes. It is found at altitudes from 1100 to
3870 m.
This species is widely distributed from central
China to the Kola Peninsula of Russia via Siberia
and Central Asia. The species is of two allopatricsubspecies, with the typical subspecies distributed
northwest of the Gobi Desert, while subspecies veitchii is in China southeast of the Gobi Desert (Fig.
3).
1a. Paeonia anomala subsp. anomala
Paeonia sinjiangensis K. Y. Pan, Fl. Reipubl. PopularisSin. 27: 603, fig. 12. 1979. TYPE: China. Xinjiang:
Habahe County, Larix forests, 1973, Y. R. Ling 1141
(holotype, PE!).
Paeonia altaica K. M. Dai & T. H. Ying, Bull. Bot. Res.
(Harbin) 10(4): 33, fig. 1. 1990. TYPE: China. Xinjiang: Habahe County, Wuzliti, 1550 m, in forests,
10 June 1980, T. H. Ying 1007 (holotype, SHMU;
isotype, PE!).

The typical subspecies is usually found in deciduous or conifer forests, in valleys, less frequently in meadows, at altitudes from 1100 to 2200 m.
Apparently it prefers relatively moist habitats. This
subspecies is of very wide distribution: northeast
Kazakhstan, northern Mongolia, from the Altai andBaikal to the Kola Peninsula in Russia, and in the
Altai and adjacent regions in Xinjiang, China (Fig.
3).
Additional specimens examined. CHINA. Xinjiang:
Altay, L. R. Xu s.n. (PE); Mt. Halamaryi, D. Y. Hong etal. Population No. 3 (PE), 0128 (PE); Xiaodong Gou valley, T. H. Ying 1001 (PE, SHMU). Burqin: Kem, N. R.
Cui 86624 (XJNU). Emin: Shiyue Township, Wuerkehe’er,

Figure 1. Paeonia holotype specimens: —A. P. anomala (LINN). —B. P. hybrida Pallas (BM). —C. P. intermedia C. A. Meyer (K): the insert at upper right is the enlargementof the label at lower right.

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Figure 2. Diagnostic features of Paeonia anomala (A, B) and P. intermedia (C, D). Tuberous or fusiform roots arecorrelated with non-caudate sepals, while carrot-shaped roots correlated with usually caudate sepals.
right bank of Telikete River, Xinjiang Branch Acad. Sinica
012 (XJBI). Fuhai: Ahebai, glades, s. coll. 3867 (XJBI);
Daqiao Forest Farm, T. H. Ying 1011 (PE, SHMU). Fuyun:
C. W. Chang 11356 (XJBI). Habahe: Baihaba, Z. M. Mao
10413 (XJBI); Tielieke, Zamanashi Valley, C. W. Changet al. 10255 (XJBI); Tiekeliti, T. H. Ying 1010 (PE,SHMU); Wuzlity, T. H. Ying 1022 (PE, SHMU). Jeminay:
Xingfu Township, Sanxie Valley, Integ. Exped. 7305
(XJBI). Qinghe: Daqinghe, Mudeng Valley, Integr. Exped.
11829 (XJBI). Tacheng: Mt. Ba’erleike, X. Y. Li 870147(SHI). Toli: Jiayi’er Gold Mine, N. R. Cui 091 (XJNU);
Mt. Zayier, N slopes, Integr. Exped. (Biology) 630 (XJBI).
Wenquan: A’erxiaoto Gou, F. M. Zhang 90-346 (XJNU).
Yumin: Ku’erzhai, Liu & Shen 8308 (XJBI). KAZAKHSTAN. Semipalatinsk Province: Altai, near Altaiskaya
Station, Mt. Narymsky region, gorge slopes, Ladygin 92(LE). Southern Altai: Mt. Ivanovsky Range, near GornoUlbinki, I. Rohderi s.n. (LE). Ustkamenogorsk region: Mt.
Altai, Kunduzda River valley, V. & A. Reznichenko 360(LE); Mt. Kalbinskie, Saryshbay River valley, V. Reznichenko 49 (LE); Mt. Kalbinskie, Ulansky, near Asybulak,
by Tainta River, P. Yurchenko s.n. (LE); Mt. Narymsky,
upper Urkar River, V. Reznichenko 58 (LE); near Ulbinsky
gorge, V. Iljin s.n. (LE); Urunhaika River mouth, S banksof Marka-Kul Lake, A. Sedelnikov s.n. (LE). Zaisan region:
Mt. Altai, Ulkunchumek River valley, upper Kaldzhir River, V. Reznichenko 106 (LE); Markakul Lake, B. Keller s.n.

(LE); Mt. Narymsky Range, Ak-tui River valley, D. Yakovlev 69 (LE); Mt. Saur, Chagan-obo River valley, V. Reznichenko s.n. (LE); Mt. Saurkie, Maly Dgewen River valley, B. Keller s.n. (LE). MONGOLIA. Ulan Bator: 10 mi.
E of Ulan Bator, P. Morgan 23 (K). Gloucester: Mongolia–Chinese Turkestan Exped. M. P. Price 293 (K). North
Mongolia: Mt. Saldzharsky Pass, E. Klements 11a (LE).
RUSSIA. Altai Region: Altai: between Charga & Shabolina, mtn. slopes, E. K. Klements 4d (LE); Chuja road,
Saadak-lary, Vereshagin 296 (LE). West Altai: Mt. Listvennitchny, near Nizhne-Uimonskoye, S. Kolomoitseva
236 (LE). Buryato-Mongolian Autonomous State:
Baikal, Tunkam, Turczaninow s.n. (LE); Zabaikalje, upper
Toreika River, 130 km W of Troitskosavsk, P. Mihno s.n.
(LE). Eubinsk Region: West Sayan, Mt. Alan Range, Golets Taskalyk, I. Krasnoborov 6765 (LE); near Buiba Station, by river, Krasnoborov & Sannikova s.n. (LE). Tuva:
Tandinsky distr., East Mt. Tannudla Range, near Shurmak,
V. Hanminchun 515 (LE); Ulug-Hemsky distr., Uiuksky
Mtn. Range, Bajan-Kol River valley, Orto-Hem tributary,
Lomonosova & Ivanova 89 (LE). Irkutsk Region: Irkutsk
Prov.: Lena River valley, upper Gulm River, I. Kuznetsov
114 (LE). Balagansky distr.: near Bazheevskoye, N. Maltsev 79 (LE). Nizhneudinsky distr.: by Angara River, opposite Yandov, S. Ganeshin s.n. (LE); Mt. Hariba-bim
Range, Angara River valley, Karpovskoye, A. Korovkin
290 (LE); by Oka River, A. Krishtofowich s.n. (LE); nearVolume 91, Number 1
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Vidimka, Vidim River, S. Kucherovskaya 282 (LE). Tunkin
distr.: Mt. Sayany, near Nylova Pustyn, V. Komarov s.n.
(LE); Nizhnaya Tunguska valley, near Nizhnaya Korelina,
S. Kokulin 254 (LE). Tunkinsky distr.: Kuren Arshan, V.
Smirnov s.n. (LE). Verholensky distr.: Biriulka, near Yushina, P. Alexandrov 179 (LE); Harat, G. Dolenko 108(LE); Lena River & Kirenga valley, Toptykan, M. Tomin
67 (LE); Tutura, P. Alexandrov 219 (LE). Kola Peninsula: Ponoi River, left bank, 8 km from its mouth, E.
Chernov 3 (LE); Ponoi River, near its mouth, N. Orlova
240 (LE). Krasnoyarsk Region: Belsk distr.: near Alshat, I. Kuznetsov 4661 (LE); Chadobets, G. Borowikow s.n.
(LE); Rybnoe, Dranitsyn & Kochubei s.n. (LE). Enisei
Prov., Achinsk distr.: near Ingol Lake, A. Suhareva s.n.
(LE); near Maloye Lake, M. Ermolajeva s.n. (LE); BolsheUluiskaya, near Bobrovki, I. Kuznetsov 53 (LE); ChukshaRiver Valley, I. Tolmatschew 247 (LE); Katanga, G. Borovikov s.n. (LE); Vyezzhi Log, A. Tugarinov s.n. (LE).
Kansk distr.: Gutar River valley, by Kamenka tributary,
W. Troitsky s.n. (LE); near Kansk, A. Shliahtin s.n. (LE);Rybnaya River valley, Perovskoye, I. Kuznetsov 193 (LE);near Nazimovskoye, Z. Evseeva 4406 (LE). Krasnoyarsk
Prov.: Chunia River, right bank opposite mouth of Mutorai
River, A. Rubin s.n. (LE); Hakassia, Shirinsky distr. Bolshaya Iiusa River valley, NW slopes, Polozhii & Kandasova s.n. (LE); near Krasnoyarsk, by Mohovaya River, V.
Verhovskaya & M. Mishin 104 (LE); Minusinsk, Bolshoye
Kysylkul Lake, K. Golubeva et al. s.n. (LE); Nizhnaya Tunguska River valley, near mouth of Hurkakit River, Rubin& Maskil s.n. (LE); between Salba & Grigorievka, P. Krylov s.n. (LE); Shushensky distr. near Tanzyben, by Black
Tanzyben River, Kuminova & Alexeeva s.n. (LE); near Sorokino Station, N slopes, Cherepnin s.n. (LE). Novosiberia Region: Altai: Elikmanar, upper Karakol River, V. Vereshagin 345 (LE). Kuznetsky Alatau: Kondoma Rivervalley, Kazany Mine, Mt. Kyon, B. Klopotov s.n. (LE); upper Sary-Chumysh River, near Ulus Munai, A. Vydrin s.n.
(LE); Tom River valley, Mt. Tohpan-taiga, B. Klopotov s.n.
(LE). Mariinsky Prov.: Biisk, Black Anuj, Talitskiebelky,
to Kazanda, Pobedimova 697 (LE); Biisk, Lebed Rivervalley, by Chuja River, B. Klopotov 180 (LE); Biisk, between Tulatinskaya & Sentelek, N. Kuznetsov 1991 (LE);Kainsk, Verhnyaya Nazarova, B. Klopotov 113 (LE); Narym, Ixa River, S. Genina s.n. (LE); Sofronova, P. Krylov
s.n. (LE); Tomsk distr.: Chulym & Chet River valley, P.
Sokolov 13a (LE). Moshkovsky Prov. (Distr.): Uskorniha
River valley, Kuvajev & Sabitov s.n. (LE). Toguchinsky
distr., near Mirny, Lashinsky & Ronginskaya 973 (LE).
Omsk Region: Tobolsk Province: Berezov distr., Lopsia
River, tributary of Severnaya Sosva, s. coll. s.n. (LE); Tobolsk distr., Landinka River valley, E. Vislouh 73 (LE).
Ural: Arctic Ural: Lyapin tributary, upper Manja River,
V. Sochava 100 (LE); Severnaya Sosva River valley, Lyapin
tributary, upper Khulga River, B. Gorodkov 512 (LE). Mariinsky distr., Beket River valley, Kolsonskoye, N. Kuznetsov 199 (LE). Tobolsk Province: Berezovsky distr., Berezov, Pohle & Rozhdestvensky s.n. (LE); Tarsky distr.,
Rybinsk, near Pustynnoye, V. Varentsov s.n. (LE). Yakutsk
Autonomous State: Yakutia, Suntarsky distr., upper Vilyuy River, Tuoikhaia, by Chona River, I. Kildushewsky
150/22 (LE).

1b. Paeonia anomala subsp. veitchii (Lynch) D.
Y. Hong & K. Y. Pan, Novon 11: 317. 2001.
Paeonia veitchii Lynch, Gard. Chron. ser. 3,
46: 2 tab. l. 1909. TYPE: China. W. Sichuan:

Hong & Pan
Paeonia anomala Complex

93

Tatien-lu (Kangding), 8–10,000 ft., E. H. Wilson (for James Veitch & Sons) 3034 (holotype,K!, photo, PE!).
Paeonia beresowskii Komarov, Bot. Mater. Gerb. Glavn.
Bot. Sada RSFSR 2: 5. 15 Jan. 1921. Paeonia veitchii var. beresowskii (Komarov) Schipczinsky, Bot.
Mater. Gerb. Glavn. Bot. Sada RSFSR 2: 46. 26 Mar.
1921. TYPE: ‘‘China Occid, Szechuan (Sichuan),Sun-pan-tin (Songpan), Guichua,’’ 9 June 1894, M.
Beresowskii s.n. (lectotype, designated here, LE notseen).
Paeonia veitchii var. woodwardii (Stapf ex Cox) Stern, J.
Roy. Hort. Soc. 68: 130. 1943. Paeonia woodwardii
Stapf ex Cox, Pl. Introd. Farrer 43. 1930. TYPE:
China. Gansu: Zone (Chuoni), R. Farrer 67 (holotype, E not seen).
Paeonia veitchii var. leiocarpa W. T. Wang & S. H. Wang,
Fl. Reipubl. Popularis Sin. 27: 603. 1979. TYPE:
China. Sichuan: Jinchuan, Kasa Township, Yinchanggou, forests by stream, 2700 m, 26 Apr. 1958,
X. Li 77248 (holotype, PE!).
Paeonia veitchii var. uniflora K. Y. Pan, Fl. Reipubl. Popularis Sin. 27: 603. 1979. TYPE: China. Sichuan:Gaze, Xiongjiling, 3600 m, mtn. summit, bushes, 18
June 1974, Qinghai-Xizang Exped. Veget. Group 034(holotype, PE!).

This subspecies, though distantly isolated from
Paeonia anomala subsp. anomala by the huge deserts of the Gobi, still very much resembles thelatter. The only remarkable difference between
them is the number of flowers on a stem. Paeonia
anomala subsp. veitchii usually possesses 2 to 4
blooming flowers in addition to up to 2 underdeveloped flower buds. Very rarely only the terminal
flower blooms in addition to up to 3 flower buds.
Paeonia anomala subsp. anomala possesses only asingle terminal blooming flower, without or infrequently with 1 to 3 underdeveloped flower buds.
The subspecies is widely distributed in China:
southeastern and central Gansu, southern Ningxia,
eastern Qinghai, the Qinling Range of Shaanxi,
Shanxi, western Sichuan, and the eastern extremeof Xizang (Tibet) (Fig. 3). Like subspecies anomala, Paeonia anomala subsp. veitchii prefers relatively moist habitats, growing in forests, grasses on
the edges of forests, bushes, or subalpine and alpine meadows with shrubs, at altitudes from 1800
to 3870 m.
Phenology. The subspecies flowers from late
April to early June, and fruits in August and September.
Chromosomes. 2n 5 10 (Hong et al., 1988).
Additional specimens examined. CHINA. Gansu: Dingxi, s. coll. s.n. (CPB). Hezheng: Xinzuang, Gansu Herbs
Group s.n. (NWTC). Hezuo: Mt. Dalinke, Q. R. Wang 7034
(NWTC). Jiangcha: S. Gansu Grassland Exped. 681(NWTC). Kangle: Mt. Taizi, G. L. Zhu 80006 (NWTC);
Mt. Lianhua, M. S. Yan 1843 (NWTC). Kangxian: s. loco,

94

Annals of theMissouri Botanical Garden

Figure 3. Distribution map of the Paeonia anomala species complex. Black circles: Paeonia anomala subsp.
anomala; white circles: P. anomala subsp. veitchii; stars: P. intermedia. Each site in the map is supported by a single
or several vouchers cited in the text.

s. coll. s.n. (NWTC). Lanzhou: Mt. Xinlong, s. coll. 84(NWTC); Mt. Tiandu, s. coll. s.n. (NWTC). Linxia: near
Senanpa, by river, T. K. Fu 834 (PE). Longdie: Dacaopo,
H. J. Zhou 708 (NWTC). Minxian: Lujing, Zhongchuangou, J. Q. Wang 197 (NWTC); Mt. Luodadoujidela, T. P.
Wang 15240 (PE); near Mawu, T. P. Wang 4594 (PE);between Lintao and Yuzhong, Huanghe Exped. 1689 (PE);Mt. Wutai, Taohe Exped. 3243 (PE); Lamathan, K. S. Hao
604 (PE). Taohe: J. F. Rock 12829 (PE). Tsaluku to Shimen, J. F. Rock 13127 (PE). Tianshui: Mt. Xinjia, W. Y.
Hsia 5707 (PE). Tianzhu: Anyuan, Y. Q. He 4516 (PE);
Zhucha, Y. Q. He 4915 (PE); Jinqiang Township, Mt. Maomao, Q. R. Wang 1869 (NWTC). Xiahe: Chingshui, T. P.
Wang 6944 (PE); Tangar’ang, Longwagou, Q. R. Wang
7270 (NWTC). Yongdeng: Liancheng, Tulugou, J. L. Bai
8613 (NWTC). Yuzhong: Xinlong, near Yangdaowan, s.
coll. s.n. (PE). Zhangxian: Caotan distr., Huanghe Exped.
03183 (PE); Mt. Guiqing, Huanghe Exped. 4735 (PE);
Hediling, Mt. Kehu, Huanghe Exped. 4655 (PE); Shichuan Forest Farm, Jinhuachi, Lian, Wang et al. 79197(NWTC). Zhouqu: Taozhou Forest Farm, Jiang & Jin
00386 (PE). Zone (Chuoni): Kache Forest Farm, Lian &Chen 31 (NWTC); Wanchang, Sangtanagou, Lian & Chen
72 (NWTC). Ningxia: the Liupan Mountain Range, Y. W.
Cui 10281 (PE). Qinghai: Datong: Kuanghuizu, K. M.
Liou 6019 (PE); near Anchiapu, K. M. Liou 5970 (PE);Dongxia, Yu, Lu, Gu & Li 68 (PE); Mt. Laoye, Liang etal. 378 (HNWP); Baoku, Z. H. Zhang et al. 4291
(HNWP). Huangyuan: Mt. Banjie, P. C. Tsoong 8842 (PE).
Huangzhong: Sanhei Forest Farm, S. X. Wang 1065
(HNWP); 40 km W of Xining, S. X. Zhen 0351 (PE).
Huzhu: Beishan Forest Farm, Guo & He 9012 (HNWP);
Qiaotou, Guo & Wang 6717 (HNWP). Jainca: near Angla,
Liu & Luo 1040 (HNWP). Ledu: Maying Township, Kongjiazui, B. Z. Guo 6767 (HNWP). Golog: Markog Forest
Farm, Hongjungou, W. Y. Wang 26818 (HNWP). Menyuan:
Semnyi, betw. Lihua & Dalong, Z. Y. Qing 1218 (HNWP,PE); Semnyi, Zhugusi, Xielong, B. Z. Guo 7396 (HNWP);
Semnyi, Hankegou, Gansu-Qinghai Exped. 2493 (PE).
Minhe: Gushan, Nanxia, L. H. Zhou 2528 (HNWP);
Xin’erbeishan, B. Z. Guo 7010 (HNWP). Tongren: Rongwo, B. Z. Guo 10232 (HNWP); Shuangfengxi, B. Z. Guo
10186 (HNWP). Xining [Sining]: Shangwuchuang, K. S.
Hao 779 (PE). Xunhua: Mengda Forest Farm, Guo &Wang 25058 (HNWP); Mt. Dalijia, G. Z. Zhang 005(HNWP). Shaanxi: Huxian: Taipingyu, Hualingwan,
Pharmac. Exped. 2019 (PE). Hwainhsien [Huayin]: Qinglongbei, W. Y. Hsia 4539 (PE). Mt. Taibai: near Dadian
temple, W. Y. Hsia 4571 (PE); Sanchaixia, K. J. Fu 4441
(PE); Dadian temple to Dumugong temple, Hong & Zhu
PB85065 (PE). Shanxi: Shiziping, Xigou, Shanxi Exped.
Wang & Tian 594 (PE). Mt. Wutai: Xitai, Xiachantan village, Xiejiagou, Shanxi Exped. 684 (PE). Sichuan: Baoxing: Raozhi, Nibagou, Hong & Zhong PB82105 (PE);Volume 91, Number 1
2004

Longdong distr., Zhonggan, Mt. Daping, Sichuan-Econom.
A59-0246 (PE); Ganyanggou, T. T. Yu 2200 (PE); Dengchigou, T. H. Tu 4329 (PE); Lianghekou, Mahuangpo, Zhang& Ren 5572 (PE). Barkam: near Puyajiao, X. Li 71029(PE); near Kanzhugou, by river, X. Li 70441 (PE);Pu’ermagou, 209 Lumbering Ground, X. Li 70867 (PE);Bu’ermagou, X. Li 70903 (PE); near Matan, X. Li 71645(PE); Dalongjiaogou, X. Li 70685 (PE); near Kafeiqiao, X.
Li 71196 (PE); Zonggag, X. Li 70719 (PE); Nazhugou, X.
Li 71087 (PE). Emei: Jieyindian, T. H. Tu 362 (PE). Dege:
S. X. Jia 229 (PE). Garze: S bank of Yalongjia River, Y.
W. Cui 4348 (PE). Heishui: Shidiaolou Township, Sichuan-Econom. A59-1208 (PE); Sandougou, Sichuan-Econom. A59-1001 (PE). Jinchuan: Zosijia, P. X. Li 10137(PE). Jiulong: Mt. Baitai, between Niupo & Baitai village,
J. S. Ying 3869 (PE). Kangding: Yulingong, Yingbapo, Hu& He 10463 (PE); Yulin Township, Simaqiao, Jiang &Xiong 35625 (PE); Yulin Township, Laoyulin, X. C. Jiang
36014 (PE); Taiyangshan distr., Hu & He 10373 (PE);Niugu village, W. Sichuan Exped. Kuan et al. 243 (PE);Xinduqiao, W. Sichuan Exped. Kuan et al. 732 (PE);Shade distr., Mi’Along, Nanshui-Beidiao Exped. 02912(PE); Zhonggu village, Dagaigou, W. Sichuan Exped. Kuan
et al. 352 (PE); between Kangding & Dawu, P. C. Tsoong
5004 (PE). Lixian: Longxi, Yiduo village, Z. He 12339(PE); Miyaluo, Misugou, S. Jiang A-7162 (PE); Shuanjingsi, P. X. Li 10100 (PE); Shuanjingsi, 333 km milestoneof Chengdu–Aba highway, Zhang & Zhou 23742 (PE);Somang, 42 km milestone of Shuanjingsi–Barkam highway, Zhang & Zhou 22675 (PE). Luhuo: P. C. Tsoong
5024 (PE); Zhehor, by river, J. S. Ying 4582 (PE). Mabian:F. T. Wang 22930 (PE). Maowen: Mt. Mafang, Sichuan
Econom. (A-59) 2554 (PE); Xiang-Naha, Sanlonggou, He& Zhou 13338 (PE). Mianning: Yejin Township, S. F. Zhu
20491(PE); Huning distr., Jiexingou, S. Jiang 5768 (PE).
Muli: Wa-chin, T. T. Yu 6137 (PE). Qianning: S of town,
Jiang & Jin 2113 (PE); to Dawu, Zhang & Lang 57 (PE).
Sêrtar: 1 km S of Wungda, S. Jiang 9082 (PE). Songpan:
Housigou, S. Jiang A-7275 (PE); Mont. Occid. H. Smith
2499 (PE). Tianquan: Mt. Erlong, Xiaoniujingou, around
Xiashan, Hu & He 10088 (PE). Wenchuan: Wolong Township, Sichuan-Econom. A59-2290 (PE); Wolong Nature
Reserve, Mt. Balang, Hong & Zhu PB85019 (PE). Xiaojin: Xihekou, Niuchanggou, Zhang & Ren 6322 (PE);Luobigou, Yangpingkou, Zhang & Ren 6082 (PE). Yajiang: Mt. Ermin, Sichuan Plants Collection 0905 (PE);Tanjiao to Malangcuo, J. S. Ying 3113 (PE). Zamtang:
Lukesi Temple, right bank, S. Jiang 8930 (PE); 10 km W
of Gangmuda, S. Jiang 8965 (PE). Xizang (Tibet): Jomda:
Gamtog distr., Qinghai-Xizang Exped, Vegetation Group
9893 (PE).

2. Paeonia intermedia C. A. Meyer, in Ledebour, Fl. Altaic 2: 277. 1830. Paeonia anomala var. hybrida f. intermedia (C. A. Meyer)
Trautvetter, Enum. Pl. Songor. 88. 1860.
Paeonia hybrida var. intermedia (C. A. Meyer)
Krylov, Fl. Altaya 1: 47. 1901. Paeonia anomala subsp. intermedia (C. A. Meyer) Trautvetter, in B. Fedtschenko, Trudy Imp. S. Petersburg. Bot. Sada 23: 351. 1904. Paeonia
anomala var. intermedia (C. A. Meyer) O. &B. Fedtsch., Beih. Bot. Centralbl. 18: 216.

Hong & Pan
Paeonia anomala Complex

95

1905. TYPE: the Altai Mountains, Ledebours.n. (lectotype, designated here, K!).
Paeonia majko N. Ketzchoweli, Notul. Syst. Geogr. Inst.
Bot. Tiphlis. fasc. 21: 17. 1959. TYPE: Georgia.
Kartly, between Igoeti and Lamitskali, 16 May 1958,
N. Ketschoweli s.n. (holotype, TBI!).
Paeonia intermedia subsp. pamiroalaica P. N. Ovchinnikov, in Fl. Tadzhikistanskoi SSR. 4: 6–10 & 531.
1975. TYPE: Tadzhikistan: declivitas australis jugi
Hissarcic, ad ripam dextram fl. Maichura, 5 km ab
ostio, 2500 m, 12.06.1971, A. A. Glebowa 6 (holotype, TAD not seen).

The species has long been identified as Paeonia
anomala (Pan, 1979), treated as a variety of P. hybrida (Krylov, 1901; Schipczinsky, 1921, 1937; Gamaulova, 1961), a variety of P. anomala (Fedtschenko, 1905; Stern, 1946; Pan, 1979), a
subspecies of P. anomala (Trautvetter, 1904), oreven as a form of P. anomala var. hybrida (Trautvetter, 1860). These diverse and erroneous assignments were made because: (1) the identity of P.
anomala, P. intermedia, and P. hybrida was not previously clear; (2) previous authors emphasized thetaxonomic value of indumentum on carpels; and (3)the characters of roots and calyces and their correlation were not considered. Examination of thetypes of the three taxa and extensive observations
on herbarium specimens and natural populations(Table 1) show that P. intermedia C. A. Meyer is anindependent species, distinctly differing from P.
anomala by its lateral roots tuberous to long-fusiform, sepals mostly (at least 2) rounded at apex but
not caudate. Also different from P. anomala, P. intermedia prefers relatively sunny and dry habitats.
Carpels vary from 1 to 5 in number, and from glabrous to densely tomentose. Ovchinnikov (1975)
properly treated the peony in Tadzhikistan as P.
intermedia, but his description of the new subspecies pamiroalaica is unjustifiable. His description
and our extensive observations have not revealed
any significant difference.
This species is widely distributed in northern
Xinjiang, China (south to the Tienshan), Kazakhstan, Kirghizia, Tadzhikistan, Uzbekistan, and theAltai of Russia. One small isolated population was
found in Georgia (Hong & Zhou, 2003) (Fig. 3). It
grows in grassy and shrubby slopes, meadows,
steppes, or in sparse woods, at altitudes from 900
to 3250 m.
Phenology. This species flowers from late May
to late June and fruits from August to September.
Chromosomes. 2n 5 10 (Hong et al., unpublished).
Additional specimens examined. CHINA. Xinjiang:
Altai Mt.: Acad. Sinica Xinjiang Integr. Exped. 10657

96

Annals of theMissouri Botanical Garden(PE). Altay: Qiao’ati, G. L. Zhu et al. 6325 (PE); to Halamaryi, Acad. Sinica Xijiang Exped. 10227 (PE); Klasu
Gou, R. C. Ching 2433 (PE); Haxiong Gou, T. Y. Chou
652100 (XJBI); Mt. Halamaryi, D. Y. Hong et al. Population No. 2 (PE); Xiaodong Gou, D. Y. Hong et al. Population No. 4 (PE). Barkol: Nanshan, s. coll. 780 (XJBI).
Fuhai: Fuhai Forest Farm, G. L. Zhu 5665 (PE); N slopes,
T. Y. Chou et al. 652150 (PE, XJBI). Fukang: Tienci, Shimen, D. Y. Hong et al. 0190 (PE). Fuyun: Mica No. 3
Mine, Integr. Exped. s.n. (XJBI); Mica No. 4 Mine, Integr.
Exped. s.n. (XJBI). Habahe: Tielieke, Kelimu 10171(XJBI); Wuzliti, Kelimu 10614 (XJBI). Hoboksar: Songshu
Gou s. coll. 7385 (XJBI); Mt. Qiaganebo, Integr. Exped.
10563 (PE, XJBI). Huocheng: Aksu Commune, Y. R. Ling
74848 (PE); Daxigou, Acad. Sinica Exped. 10402 (PE);Xin’ertai, Yeguolin, C. W. Chang et al. 3316 (XJBI);
Xiaoxigou, A. R. Li & J. N. Zhu 10414 (PE, XJBI). Mori:
Nangou, S. L. Chen 0040 (PE). Qapqal: 58 km from Yining
to Zhaosu, Inst. Biology and Pedology 66 (XJBI). Qitai:
Biliuhe, J. C. Zhao 82-4288 (PE, XJU); Kuangou, Biology
81-8381574 (XJBI); Tangfangmen Forest Farm, Integr.
Inst. Bio. Res. 5788 (XJBI); Yuejin Commune, S. Q. Lin
et al. 34 (XJBI). Tacheng: Mt. Baerleike, N slope, Integ.
Exped. 1200 (XJBI). Toli: Mt. Albakzin, mountain slope,
K. C. Kuan 2611 (PE, XJBI). Xinyuan: Nanshan, T. Y.
Chou et al. 650392 (PE). Yining: Yining Forest Farm, Jirtoulong valley, D. Y. Hong et al. Population 1 & 092 (PE).
Yumin: The Wild Bada Prunus Nature Reserve, J. C. Zhao
85-293 (PE, XJU). GEORGIA. Kartly: between Lamistskali and Igoeti, Hong & Zhou H99033 (PE). KAZAKHSTAN. Ili-Pribalkhash: NW ridges of Mt. Kirgizsky
Range, upper Sugata River, V. Golos-Kokov s.n. (LE); Issyk-Kul Province, Santash, Dongulsky garden, Aidarova& Kashenko s.n. (LE); Semirechensk Province, N slope of
Alatau, between Verny to Kastek mountain pass, V. Ladygin s.n. (LE); SW Songorian Alatau, Mt. Emelsky Range,
Mt. Matai, V. Goloskokov s.n. (LE); Taldy-Kurgan Province,
Mt. Altyn-Emel Range, Mt. Engi-Altyn-Emel pass to roadSary-Ozek-Dzharkent, Grubov & Luboarsky 229 (LE); Zailiisky Alatau, Usun-Kargali River Valley, W Sukonnaya
Fabrika, Y. Goloskokov (LE). E. Kazakhstan: Mt. Kalbinsky Range, Mt. Kok-tau, near Sabinskie Lake, A. Yunatov
24 (LE); Semipalatinsk Province, Zaisan distr., Kusto
Gorge, B. Shishkin s.n. (LE); Zaisan distr., near Kenderlyk, B. Shishkin s.n. (LE); near Zaisan, Mt. Kichken-tau,
Tatarowa & Borsowa 52 (LE). Mountain Kazakhstan:
Semirechensk Prov., Songorian Alatau, S of Lepsinsk, R.
Rozhevits 55 (LE); W ridges of Songorian Alatau, Koksu
Gorge, near Koksuisky, V. Goloskokov (LE); Tarbagatai,
Chenarak, steppes, Grigoriev & Buhaeva 298 (LE); Mt.
Tsarytsynsky Pass, deep gorge from the pass, Schipczinsky
125 (LE). S. Kazakhstan: Mt. Chatkalsky Range, Chatkal Valley, Mt. Piazak, O. Knorring 60 (LE); Prichuisky
Mujun(Kum), Sulunger Gorge, F. Zapriagaev 66 (LE).
KIRGHIZIA. Dzhalalabad Province: Bazar-Kurgan
distr., Mt. Fergansky Range, S slopes, near AK-Terek, Lebedeva & Faleeva s.n. (LE); Kara-Alma, Toguz (Bulak), N
slopes, Poniatovskaya, s.n. (LE); Tianshan, Mt. Ketmen
Range, Mt. Elchin-Ustriuk, Kizilbulak Gorge, s. coll. s.n.
(LE). Fergana Province: Kirgizsky Alatau, Makbal
Gorge, Ty-Karyn River, G. Igolkin 97 (LE); Oshsky distr.,
Ak-Bura valley, Mt. Kanida Pass, O. Knorring 25 (LE).
Semirechensk Province: Pishpeksky distr., Uzungyrskaya Volost, Mt. Aleksandrovsky Range, Mt. Shekule N
slope, A. Savenkova 29 (LE). RUSSIA. Altai: Alex Shrenks.n. (LE); Altai, Barnaul Province, between Kalmytskie
Mysy & Mahanova, P. Krylov s.n. (LE). TADZHIKISTAN.

Gissarsky Mountain Range: Buhara, Gissar, Mtn. Zarkamar Pass, V. Lipsky 1697 (LE). Darvazsky Mtn. Range,
N slopes, left bank of Zarako River, opposite Havdak,
Shinova & Kinzikaeva 1550 (LE). Imam-Askari, Mt. W of
Darai-Imam, Bochantsev & Egorova 1014 (LE). Peter I
Range, N slopes, Nazarak Valley, upper reaches, 15 km S
of Tadzhikabad, Ladygina, Ikonnikov & Fridman 1337
(LE). Vahshsky Mtn. Range, Mt. Hozreshi-Sho, above
Sary-Hosor, Sidorenko 185 (LE). Varzob Valley, right bankof Maihur Gorge, NE slope, Stepanenko & Kudriashova
2514 (LE); Mountain pass between rivers Havag & Ketsgug, N. Gontscharow 2051 (LE); upper Horanchon, Zapriagaev 351 (LE). UZBEKISTAN. Tashkent Province:
Bolshoy Chimgan, Z. von Minkwitz 1190 (LE); Bolshoy
Chimgan, N stony slopes, V. Bochantsev 547 (LE).
Literature Cited
Anderson, G. 1818. A monograph of the genus Paeonia.
Trans. Linn. Soc. London. 12(1): 248–283.
Dai, K. M & T. H. Ying. 1990. A new species of the genus
Paeonia from China. Bull. Bot. Res. (Harbin) 10(4): 33–
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URSS 7: 24–35. Editio Akademiae Scientiarum URSS,
Leningrad.
Schmitt, E. 1999. Les Pivoines. Étude systematique dugenre Paeonia L. Plantes de Montagne 191: 574–583.
Stern, F. C. 1946. A Study of the Genus Paeonia. Royal
Horticultural Society, London.
Trautvetter, E. R. 1860. Enumeratio plantarum songori-Volume 91, Number 1
2004carum, a Dr. Alex Schrenk annis 1840–1843 collectarum. Bull. Soc. Nat. Mosc. 33: 87–88.
. 1904. Paeonia. In: B. A. Fedtschenko (editor),
Flora of the West Tianshan (Acta Hort. Petrop.) 23:
351–353.
APPENDIX 1
INDEX TO EXSICCATAE EXAMINEDThe figures in parentheses represent: 1a 5 Paeonia anomala subsp. anomala; 1b 5 P. anomala subsp. veitchii;
2 5 P. intermedia.
Acad. Sinica Bot. Inst. Exped. Xinjiang 1803 (PE,XJBI) (2); Acad. Sinica Exped. 10402 (PE) (2); Acad. Sinica Xinjiang Exped. 2496 (PE, XJBI) (1a), 10227 (PE)(2); Acad. Sinica Xinjiang Integr. Exped. 10657 (PE) (2);Agr. College 1051 (PE) (2); Aidarova & Kashenko s.n.
(LE) (2); Alexandrov, P. 179 (LE) (1a), 219 (LE) (1a); Alex
Shrenk s.n. (LE) (2).
Bai, J. L. 8613 (NWTC) (1b); Beresowskii, M. s.n. (LE)(1b, type of Paeonia beresowskii Komarov); Biology Exped.
79250198 (XJBI) (2), 81-8330478 (XJBI) (2); Bochantsev,
V. 547 (LE) (2); Bochantsev & Egorova 1014 (LE) (2);
Borovikov, G. s.n. (LE) (1a); Borovikov et al. s.n. (LE) (1a).
Chang, C. W. 11356 (XJBI) (1a); Chang, C. W. et al.
3316 (XJBI) (2), 4037 (XJBI) (2), 4357 (XJBI) (1a), 10255(XJBI) (1a); Chen & Ju s.n. (NWTC) (1b); Chen, S. L.
0040 (PE) (2); Cherepnin s.n. (LE) (1a); Ching, R. C. 1803
(PE) (1a), 2433 (PE) (2), 2496 (PE) (1a); Chou, T. Y.
652100 (XJBI) (2); Chou, T. Y. et al. 650392 (PE) (2),650868 (PE, XJBI) (2), 652150 (PE, XJBI) (2); Cui, N.
R. 091 (XJNU) (1a), 86624 (XJNU) (1a); Cui, Y. W. 4348
(PE) (1b), 10281 (PE) (1b).
Dolenko, G. 108 (LE) (1a); Dranitsyn & Kochubei s.n.
(LE) (1a).
Ermolajeva, M. s.n. (LE) (1a); Evseeva, Z. 4406 (LE)
(1a).
Fang, W. P. 4213 (LE) (1a), 6037 (LE) (1a); Farrer, R.
67 (E) (1b, type of Paeonia woodwardii Stapf ex Cox);
Fedorov & Iljina 72 (LE) (2), 128 (LE) (2); Fu, K. J. 4441
(PE) (1b); Fu, T. K. 834 (PE) (1b).
Ganeshin, S. s.n. (LE) (1a); Gansu Herbs Group s.n.
(NWTC) (1b); Genina, S. s.n. (LE) (1a); Gansu-Qinghai
Exped. 2493 (PE) (1b); Goloskokov, V. s.n. (LE) (2); Gontscharow, N. 2051 (LE) (2); Gorodkov, B. 512 (LE) (1a);Grigoriev & Buhaeva 298 (LE) (2); Grubov & Luboarsky
229 (LE) (2); Guo, B. Z. 6767 (HNWP) (1b), 7010
(HNWP) (1b), 7396 (HNWP) (1b), 10186 (HNWP) (1b),10232 (HNWP) (1b); Guo & He 9012 (HNWP) (1b); Guo& Wang 6008 (HNWP) (1b), 6717 (HNWP) (1b), 25058(HNWP) (1b); Guo & Yang 9608 (HNWP) (1b).
Hanminchun, V. 515 (LE) (1a); Hao, K. S. 779 (PE)(1b); He, Y. Q. 4516 (PE) (1b), 4915 (PE) (1b); He, Z.
12339 (PE) (1b); He & Zhou 13338 (PE) (1b); Hong D.
Y. et al. Population 2 (PE) (2), Population 3 (PE) (1a),Population 4 (PE) (2), Population 5 (PE) (1a), Population
1 (PE) (2), Population 0190 (PE) (2); Hong D. Y. & Zhong
PB82105 (PE) (1b); Hong D. Y. & Zhu PB85019 (PE)(1b), PB85040 (PE) (1b), PB85065 (PE) (1b); Hong, Luo& He H95034 (A, CAS, K, MO, PE, US) (1b); Hong &Zhou H99033 (PE) (2); Hsia, W. Y. 4539 (PE) (1b),4571(PE) (1b), 5707 (PE) (1b); Hu, W. G. 13050 (PE)(1b); Huanghe Exped. 1689 (PE) (1b), 3183 (PE) (1b),4655 (PE) (1b), 4735 (PE) (1b); Hu & He 10088 (PE)(1b), 10373 (PE) (1b), 10463 (PE) (1b), 11173 (PE) (1b);Huang, Luo & Jiang 730 (PE) (1b).
Igolkin, G. 97 (LE) (2); Ikonnikov & Fridman 892 (LE)Hong & Pan
Paeonia anomala Complex

97(2); Iljin, V. s.n. (LE) (1a); Integr. Exped. 630 (XJBI) (1a),1200 (XJBI) (2), 7305 (XJBI) (1a), 10563 (PE, XJBI) (2),11829 (XJBI) (1a), s.n. (XJBI) (2); Integr. Inst. Hydrol.
Pedol & Biol. Res. 5788 (XJBI) (2).
Jia, S. X. 229 (PE) (1b); Jiang & Jin 00386 (PE) (1b),2113 (PE) (1b); Jiang & Xiong 34242 (PE) (1b), 34319(PE) (1b), 35625 (PE) (1b); Jiang, S. 5768 (PE) (1b), 8874
(PE) (1b), 8930 (PE) (1b), 8965 (PE) (1b), 9082 (PE) (1b),A-7162 (PE) (1b), A-7275 (PE) (1b); Jiang, X. C. 36014(PE) (1b), 36433 (PE) (1b).
Kelimu 10171 (XJBI) (2), 10369 (XJBI) (2), 10614(XJBI) (2); Keller, B. s.n. (LE) (1a); Ketzchoweli, N. s.n.
(TBI) (2, type of Paeonia majko N. Ketzchoweli); Kildushewsky, I. 150/22 (LE) (1a); Klements, E. 4d (LE) (1a),11a (LE) (1a); Klopotov, B. 113 (LE) (1a), 180 (LE) (1a);Knorring, O. 25 (LE) (2), 66 (LE) (2); Kokulin, S. 254(LE) (1a); Kolomoitseva, S. 236 (LE) (1a); Komarov, V. s.n.
(LE) (1a); Korovkin, A. 290 (LE) (1a); Krasnoborov, I.
6765 (LE) (1a); Krishtofowich, A. s.n. (LE) (1a); Krylov,
P. s.n. (LE) (2); Kuan, K. C. 2611 (PE, XJBI) (2), 77309(PE) (1b); Kuan et al. 243 (PE) (1b), 352 (PE) (1b); Kucherovskaya, S. 282 (LE) (1a); Kuminova & Alexeeva s.n.
(LE) (1a); Kuvajev & Sabitov s.n. (LE) (1a); Kuznetsov, I.
53 (LE) (1a), 114 (LE) (1a), 193 (LE) (1a), 4661 (LE) (1a);Kuznetsov, N. 199 (LE) (1a), 1991 (LE) (1a).
Ladygin 92 (LE) (1a), s.n. (LE) (2); Ladygina et al. 1337
(LE) (2); Lashinsky & Ronginskaya 973 (LE) (1a); Lebedeva & Faleeva s.n. (LE) (2); Ledebour, s.n. (K) (2, type ofPaeonia intermedia C. A. Meyer); Li, A. R. & Zhu, J. N.
10414 (PE, XJBI) (2); Li, P. X. 10100 (PE) (1b), 10137(PE) (1b), 10143 (PE) (1b); Li, X. 70247 (PE) (1b), 70441(PE) (1b), 70510 (PE) (1b), 70685 (PE) (1b), 70719 (PE)(1b), 70867 (PE) (1b), 70903 (PE) (1b), 71029 (PE) (1b),71087 (PE) (1b), 71196 (PE) (1b), 71645 (PE) (1b), 74842(PE) (1b), 74936 (PE) (1b), 77248 (PE) (1b, type ofPaeonia veitchii var. leiocarpa W. T. Wang & S. H. Wang);
Li, X. Y. 870147 (SHI) (1a), 870169 (SHI) (2); Lian &Chen 31 (NWTC) (1b), 72 (NWTC) (1b); Lian et al. 378(HNWP) (1b), 79179 (NWTC) (1b); Licent, P. 2022 (PE)(1b); Lin, S. Q. et al. 34 (XJBI) (2); Ling, Y. R. 1141 (PE)
(1a, type of Paeonia sinjiangensis K. Y. Pan), 74243 (PE)(2), 74848 (PE) (2), 74849 (PE) (2); Liou, K. M. 5659
(PE) (1b), 5970 (PE) (1b), 6019 (PE) (1b), 6178 (PE) (1b);
Lipsky, V. 1697 (LE) (2); Liu & Luo 1040 (HNWP) (1b);
Liu & Shen 8308 (XJBI) (1a); Liu, S. W. 2115 (HNWP)(1b); Lomonosova & Ivanova 89 (LE) (1a); Long, X. F. 85(NWTC) (1b).
Maltsev, N. 79 (LE) (1a); Mao, Z. M. 10413 (XJBI) (1a);Mihno, P. s.n. (LE) (1a); Minkwitz Z. von 1190 (LE) (2);Morgan, P. 23 (K) (1a).
Nanshui-Beidiao Exped. 02912 (PE) (1b).
Pharmac. Exped. 2019 (PE) (1b); Pobedimova 697 (LE)(1a); Pohle & Rozhdestvensky s.n. (LE) (1a); Polozhii &Kandasova s.n. (LE) (1a); Poniatovskaya s.n. (LE) (2);Price, M. P. 293 (LE) (1a).
Qing, Z. Y. 1218 (HNWP, PE) (1b); Qinghai-Xizang Exped. Veget. Group 034 (PE) (1b, type of Paeonia veitchii
var. uniflora K. Y. Pan), 9893 (PE) (1b); Qu, Z. X. 2792
(PE) (1b).
Reznichenko, V. 49 (LE) (1a), 58 (LE) (1a); Reznichenko, V. & A. 106 (LE) (1a), 360 (LE) (1a), s.n. (LE) (1a);Rock, J. F. 12260 (PE) (1b), 12829 (PE) (1b), 13127 (PE)(1b); Rohderi, I. s.n. (LE) (1a); Rozhevits, R. 55 (LE) (2);
Rubin, A. s.n. (LE) (1a); Rubin & Maskil, s.n. (LE) (1a).
Sanadze et al. s.n. (TBI) (2); Savenkova, A. 29 (LE) (2);Schipczinsky 125 (LE) (2); s. coll. Siberia, (LINN) (1a,type of Paeonia anomala Linnaeus); Sedelnikov, A. s.n.

98Annals of theMissouri Botanical Garden(LE) (1a); S. Gansu Grassland Exped. 681 (NWTC) (1b);Shanxi Exped. 684 (PE) (1b); Shanxi Exped. Wang & Tian
594 (PE) (1b); Shinova & Kinzikaeva 1550 (LE) (2);Shishkin, B. s.n. (LE) (2); Shliahtin, A. s.n. (LE) (1a);Shukin, S. s.n. (LE) (1a); Sichuan-Econom. A59-0246(PE) (1b), A59-1001 (PE) (1b), A59-1208 (PE) (1b), A592554 (PE) (1b), Sichuan-Econom. & Ya 838 (PE) (1b);Sichuan Plants Collection 0905 (PE) (1b); Sidorenko 185(LE) (2); Smirnov, V. s.n. (LE) (1a); Smith, H. 2499 (PE)(1b); Sochava, V. 100 (PE) (1b); Sokolov, P. 13a (LE) (1a);Song, Z. P. 38544 (PE) (1b), 39107 (PE) (1b); Stepanenko& Kudriashova 2514 (LE) (2); Suhareva, A. s.n. (LE) (1a).
Tang, T. 938 (PE) (1b); Taohe Exped. 3243 (PE) (1b);Tatarova & Borsowa 52 (LE) (2); Tolmatschew, I. 247 (LE)(1a); Tomin, M. 67 (LE) (1a); Tripolitova, T. s.n. (LE) (2);Troitsky, W. s.n. (LE) (1a); Tsoong, P. C. 5004 (PE) (1b),5024 (PE) (1b), 8842 (PE) (1b); Tu, T. H. 362 (PE) (1b),4329 (PE) (1b); Tugarinov, A. s.n. (LE) (1a); Turczaninow,
s.n. (LE) (1a).
Varentsov, V. s.n. (LE) (1a); Vereshagin, V. 296 (LE)(1a), 345 (LE) (1a); Verhovskaya & Mishin 104 (LE) (1a);Vislouh, E. 73 (LE) (1a); Vydrin, A. s.n. (LE) (1a).
Wang, F. T. 22930 (PE) (1b); Wang, J. Q. 197 (NWTC)(1b); Wang, Q. R. 1869 (NWTC) (1b), 7034 (NWTC) (1b),7270 (NWTC) (1b); Wang, S. X. 1065 (HNWP) (1a);Wang, T. P. 197 (NWTC) (1a), 15240 (NWTC) (1a), 4594
(NWTC) (1a), 6944 (PE) (1b); Wang, W. Y. 26771 (HNWP)(1b), 26818 (HNWP) (1b), 27060 (HNWP) (1b); Wang, W.

Y. et al. 27347 (HNWP) (1b); Wang & Zhou 19 (HNWP)(1b), 198 A (HNWP) (1b).
Xinjiang Branch Acad. Sinica 012 (XJBI (1a); Xu, L.
R. s.n. (PE) (1a).
Yakovlev, D. 69 (LE) (1a); Yan, M. S. 1843 (NWTC)(1b); Ying, J. S. 3113 (PE) (1b), 3869 (PE) (1b), 4582
(PE) (1b), 4591 (PE) (1b); Ying, T. H. 995 (PE, SHMU)(1a), 1001 (PE, SHMU) (1a), 1006 (PE, SHMU) (1a), 1007
(SHMU, PE) (1a, type of Paeonia altaica K. M. Dai & T.
H. Ying), 1008 (PE) (1a), 1009 (PE, SHMU) (1a), 1010
(PE, SHMU) (1a), 1011 (PE, SHMU) (1a), 1014 (PE,SHMU) (1a), 1022 (PE, SHMU) (1a); Yu et al. 68 (PE)(1b); Yu, T. T. 2200 (PE) (1b), 6137 (PE) (1b); Yunatov,
A. 24 (LE) (2); Yurchenko, P. s.n. (LE) (1a).
Zapriagaev, F. 66 (LE) (2), 351 (LE) (2); Zhang, F. M.
90-346 (XFNU) (1a); Zhang, G. Z. 005 (HNWP) (1b);Zhang & Lang 57 (PE) (1b); Zhang & Li 0502 (HNWP)(1b); Zhang & Ren 5572 (PE) (1b), 6082 (PE) (1b), 6322
(PE) (1b); Zhang, Z. H. et al. 0097 (HNWP) (1b), 0414
(HNWP) (1b), 4291 (HNWP) (1b), 4317 (HNWP) (1b),4320 (HNWP) (1b); Zhang & Zhou 22092 (PE) (1b),22155 (PE) (1b), 22675 (PE) (1b), 23742 (PE) (1b); Zhao,
J. C. 82-293 (PE, XJU) (2), 82-4288 (PE, XJU) (2), s.n.
(PE, XJU) (2); Zhen, S. X. 0351 (PE) (1b); Zhou, H. J.
708 (NWTC) (1a); Zhou, L. H. 2528 (HNWP) (1b); Zhu,
G. L. 5665 (PE) (2), 5755 (PE) (1a), 6345 (PE) (1a),80006 (NWTC) (1b); Zhu, G. L. et al. 6325 (PE) (2), 6386
(PE) (2); Zhu, S. F. 20491 (PE) (1b).